Black Hills Spruce
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''Picea glauca'', the white spruce, is a species of
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfami ...
native to the northern temperate and boreal forests in North America. ''Picea glauca'' is native from central Alaska all through the east, across southern/central Canada to the
Avalon Peninsula The Avalon Peninsula (french: Péninsule d'Avalon) is a large peninsula that makes up the southeast portion of the island of Newfoundland. It is in size. The peninsula is home to 270,348 people, about 52% of Newfoundland's population, according ...
in Newfoundland, and south to Montana, Minnesota, Wisconsin, Michigan, Upstate New York and Vermont, along with the mountainous and immediate coastal portions of New Hampshire and Maine, where temperatures are just barely cool and moist enough to support it. There is also an isolated population in the Black Hills of South Dakota and Wyoming. It is also known as Canadian spruce, skunk spruce, cat spruce, Black Hills spruce, western white spruce, Alberta white spruce, and Porsild spruce.


Description

The white spruce is a large evergreen
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
which normally grows to tall, but can grow up to tall with a trunk diameter of up to . The
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
is thin and scaly, flaking off in small circular plates across. The crown is narrowconical in young trees, becoming cylindrical in older trees. The shoots are pale buff-brown, glabrous in the east of the range, but often
pubescent The adjective pubescent may describe: * people or animals undergoing puberty * plants that are hairy, covered in trichomes * insects that are covered in setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a ...
in the west, and with prominent pulvini. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are needle-like, 12 to 20 millimetres long,
rhombic Rhombic may refer to: * Rhombus, a quadrilateral whose four sides all have the same length (often called a diamond) *Rhombic antenna, a broadband directional antenna most commonly used on shortwave frequencies * polyhedra formed from rhombuses, suc ...
in cross-section,
glaucous ''Glaucous'' (, ) is used to describe the pale grey or bluish-green appearance of the surfaces of some plants, as well as in the names of birds, such as the glaucous gull (''Larus hyperboreus''), glaucous-winged gull (''Larus glaucescens''), g ...
blue-green above (hence ''glauca'') with several thin lines of
stomata In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
, and blue-white below with two broad bands of stomata. The cones are pendulous, slender, cylindrical, 3 to 7 cm long and 1.5 cm wide when closed, opening to 2.5 cm broad. They have thin, flexible scales 15  mm long with a smoothly rounded margin. They are green or reddish, maturing to pale brown 4 to 8 months after pollination. The seeds are black, 2 to 3 mm long, with a slender, 5 to 8 mm long pale brown wing. File:Picea glauca sapling Kluane NP.jpg, Young tree File:2009-05-13 EPB mature (02) DDumais.jpg, Mature tree File:P1260709 DDumais.jpg, Mature tree that has lost its lower branches File:Picea glauca-bark.jpg, The
bark Bark may refer to: * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Places * Bark, Germany * Bark, Warmian-Masurian Voivodeship, Poland Arts, ...
is thin and scaly, flaking off in small circular plates File:Picea glauca twig Denali NP AK.jpg, Twig with striped, blue-green, four-sided needles File:Picea glauca Świerk biały 2011-09-11 03.jpg, Mature female cone File:Picea glauca Brno3.JPG, Female cone File:Picea glauca young female cone - Keila.jpg, Young female cone File:Picea glauca 2.jpg, Male cone and pollen


Seeds

Seeds are small, 2.5 to 5 mm long, oblong, and acute at the base. Determinations of the average number of sound seeds per white spruce cone have ranged from 32 to 130.Zasada, J.C.; Viereck, L.A. 1970. White spruce cone and seed production in interior Alaska, 1957–68. USDA, For. Serv., Pacific NW For. Range Exp. Sta., Portland OR, Res. Note PNW-129. 11 p.
oates et al. 1994 Oates may refer to: ;People * Oates (surname) *Garfunkel and Oates, comedy act ** Kate Micucci (aka ''Oates'') of "Garfunkel and Oates" *Hall & Oates, musical act ;Places and buildings * Oates, Missouri, a community in the United States *Oates Coa ...
/ref> Common causes of empty seed are lack of pollination, abortion of the ovule, and insect damage. The average weight per individual seed varies from 1.1 mg to 3.2 mg. Each seed is clasped by a thin wing 2 to 4 times as long as the seed. Seed and wing are appressed to the cone scale. Embryo and
megagametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
are soft and translucent at first; later the endosperm becomes firm and milky white, while the embryo becomes cream-coloured or light yellow. At maturity, the testa darkens rapidly from light brown to dark brown or black.Crossley, D.I. 1953. Seed maturity in white spruce. Canada Dep. Resour. and Devel., For. Branch, For. Res. Div., Ottawa ON, Silv. Res. Note 104. 16 p. Mature seeds “snaps in two” when cut by a sharp knife on a firm surface. White spruce cones reach their maximum size after 800 GDD. Cone moisture content decreases gradually after about 1000 GDD.Cram, W.H.; Worden, H.A. 1957. Maturity of white spruce cones and seed. For. Sci. 3:263–269. Cone colour also can be used to help determine the degree of maturation, but cones may be red, pink or green.Teich, A.H. 1970. Genetic control of female flower colour and random mating in white spruce. Can. Dep. Fish. For., Can. For. Serv., Ottawa ON, Bi-mo. Res. Notes 26:2. Collection and storage dates and conditions influence
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
requirements and early seedling growth.Zasada, J.C. 1973. Effect of cone storage method and collection date on Alaskan white spruce (''Picea glauca'') seed quality. p. 1–10 (paper 19) ''in'' Proc. Seed Problems. IUFRO Symp. Seed Processing, Bergen, Norway. Working Party S2.01, Royal Coll. For., Bergen, Norway, Vol. 1.
oates et al. 1994 Oates may refer to: ;People * Oates (surname) *Garfunkel and Oates, comedy act ** Kate Micucci (aka ''Oates'') of "Garfunkel and Oates" *Hall & Oates, musical act ;Places and buildings * Oates, Missouri, a community in the United States *Oates Coa ...
/ref>Edwards, I.K. 1977. Fertility of transplant fields at the Prince Albert Forest Nursery. Can. Dep. Fish. Environ., Can. For. Serv., Northern For. Res. Centre, Edmonton AB, Inf. Rep. NOR-X-189. 21 p.Winston, D.A.; Haddon, B.D. 1981. Effects of early cone collection and artificial ripening on white spruce and red pine germination. Can. J. For. Res. 11:817–826. A bushel (35 L) of cones, which may contain 6500 to 8000 cones, yields of clean seed.USDA Forest Service. 1948. Woody-plant Seed Manual. USDA, For. Serv., Washington DC, Misc. Publ. 654. 416 p. Seed dispersal begins after cone scales reflex with cone maturation in the late summer or early fall of the year of formation. Cones open at moisture contents of 45% to 70% and specific gravities of 0.6 to 0.8. Weather affects both the initiation and pattern of seed dispersal, but cone opening and the pattern of seed dispersal can vary among trees in the same stand. Even after dispersal has begun, cold, damp weather will cause cone scales to close; they will reopen during dry weather. Most seed falls early rather than late, but dispersal may continue through fall and winter and even into the next growing season.Zasada, J. 1986. Natural regeneration of trees and tall shrubs on forest sites in interior Alaska. p. 44–73'' in'' Van Cleve, K.; Chapin, F.S.; Flanagan, P.W.; Viereck, L.A.; Dyrness, C.T. (Eds.). Forest Ecosystems in the Alaskan Taiga: a Synthesis of Structure and Function. Springer-Verlag New York NY.Rowe, J.S. 1953. Viable seed on white spruce trees in midsummer. Can. Dep. Northern Affairs and National Resources, For. Branch, For. Res. Div., Ottawa ON, Silv. Leafl. 99. 2 p. Seed dispersal occurs mainly in late summer or early fall. White spruce seed is initially dispersed through the air by wind. Both the initiation and pattern of seed dispersal depend on the weather, but these can vary among trees in the same stand. Small amounts of white spruce seed are normally dispersed beyond 100 m from the seed source, but exceptionally seeds have been found more than 300–400 m from the nearest seed source.


Root system

The root system of white spruce is highly variable and adaptable, responding to a variety of edaphic factors, especially
soil moisture Soil moisture is the water content of the soil. It can be expressed in terms of volume or weight. Soil moisture measurement can be based on ''in situ'' probes (e.g., capacitance probes, neutron probes) or remote sensing methods. Water that enters ...
,
soil fertility Soil fertility refers to the ability of soil to sustain agricultural plant growth, i.e. to provide plant habitat and result in sustained and consistent yields of high quality.
, and mechanical impedance.Wagg, J.W.G. 1964. White spruce regeneration on the Peace and Slave River lowlands. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1069. 35 p.Wagg, J.W.B. 1967. Origin and development of white spruce root-forms. Can. Dep. For. Rural Devel., For. Branch, Ottawa ON, Publ. 1192. 45 p. On soils that limit rooting depth, the root system is plate-like, but it is a common misconception to assume that white spruce is genetically constrained to develop plate-like root systems irrespective of soil conditions.Sutton, R.F. 1969. Form and development of conifer root systems. Commonw. For. Bureau, Oxford, U.K., Tech. Communication No. 7. 131 p. In the nursery, or naturally in the forest, white spruce usually develops several long 'running' roots just below the ground surface.Mullin, R.E. 1957. Experiments with root and top pruning of white spruce nursery stock. Ont. Dep. Lands For., Res. Div., Toronto ON, Res. Rep. 36. 31 p. The structure of the tracheids in the long lateral roots of white spruce varies with soil nitrogen availability.Krasowski, M.J.; Owens, J.N. 1999. Tracheids in white spruce seedling’s long lateral roots in response to nitrogen availability. Plant and Soil 217(1/2):215–228.


Stem

White spruce can live for several hundred years, with an estimated average lifespan of 250 to 300 years.Dallimore, W.; Jackson, A.B. 1961. A Handbook of Coniferae including Ginkgoaceae, 3rd (1948) ed. reprinted with corrections. Arnold, London, U.K. 686 p. Slow-growing trees in rigorous climates are also capable of great longevity. White spruce high on the shore of Urquhart Lake, Northwest Territories, were found to be more than 300 years old.Hare, F.K.; Ritchie, J. 1972. The boreal bioclimates. Geogr. Rev. 62:333–365.


Bark

The bark of mature white spruce is scaly or flaky, grey-brown or ash-brown, but silvery when freshly exposed.Brayshaw, T.C. 1960. Key to the native trees of Canada. Canada Dep. For., Bull. 125. 43 p.Harlow, W.M.; Harrar, E.S. 1950. Textbook of Dendrology, 3rd ed. McGraw-Hill, New York NY. 555 p. Resin blisters are normally lacking, but the Porsild spruce ''Picea glauca'' var. ''porsildii'' Raup has been credited with having smooth resin-blistered bark.Hosie, R.C. 1969. Native Trees of Canada, 7th ed. Can. Dep. Fish. For., Can. For. Serv., Ottawa ON. 380 p. White spruce bark is mostly less than 8 mm and not more than 9.5 mm thick.Hale, J.D. 1955. Thickness and density of bark. Pulp and Paper Mag. Canada, Dec.:3–7.Chang, Y.P. 1954. Bark structure of North American conifers. USDA, For. Serv., Tech. Bull. 1095. 86 p.


Chemistry

Isorhapontin Isorhapontin is a stilbenoid. It is the glucoside of isorhapontigenin. It can be found in mycorrhizal and non-mycorrhizal roots of Norway spruces (''Picea abies''), in the bark of ''Picea sitchensis'' or in white spruce (''Picea glauca ''Picea ...
can be found in spruce species such as the white spruce.


Distribution

White spruce has a transcontinental range in North America. In Canada, its
contiguous distribution Species distribution —or species dispersion — is the manner in which a biological taxon is spatially arranged. The geographic limits of a particular taxon's distribution is its range, often represented as shaded areas on a map. Patterns of ...
encompasses virtually the whole of the
Boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
, Subalpine, Montane, Columbia, Great Lakes–St. Lawrence, and
Acadian Forest The Acadians (french: Acadiens , ) are an ethnic group descended from the French who settled in the New France colony of Acadia during the 17th and 18th centuries. Most Acadians live in the region of Acadia, as it is the region where the des ...
Regions, extending into every province and territory.Rowe, J.S 1972. Forest regions of Canada. Can. Dep. Environ., Can. For. Serv., Ottawa ON, Publ. 1300. 172 p.Forestry Branch. 1961. Native Trees of Canada, 6th ed. Canada Dep. Northern Affairs and National Resour., For. Branch, Ottawa ON, Bull. 61. 291 p. On the west coast of
Hudson Bay Hudson Bay ( crj, text=ᐐᓂᐯᒄ, translit=Wînipekw; crl, text=ᐐᓂᐹᒄ, translit=Wînipâkw; iu, text=ᑲᖏᖅᓱᐊᓗᒃ ᐃᓗᐊ, translit=Kangiqsualuk ilua or iu, text=ᑕᓯᐅᔭᕐᔪᐊᖅ, translit=Tasiujarjuaq; french: b ...
, it extends to Seal River, about 59°N, "from which the northward limit runs apparently almost directly north-west to near the mouth of the Mackenzie River, or about latitude 68°".Bell, R. 1881. The northern limits of the principal forest trees of Canada east of the Rocky Mountains. p.38c–56c ''in'' Geological and Natural History Survey of Canada, Ottawa ON, Report 1879/1880. Collins and SumnerCollins, G.L.; Sumner, L. 1953. Northeast Arctic: the last great wilderness. Sierra Club Bull. 38:13–26. reported finding white spruce within 13 km of the Arctic coast in the
Firth Valley Firth is a word in the English and Scots languages used to denote various coastal waters in the United Kingdom, predominantly within Scotland. In the Northern Isles, it more usually refers to a smaller inlet. It is linguistically cognate to ''fj ...
, Yukon, at about 69°30′ N, 139°30′ W. It reaches within 100 km of the Pacific Ocean in the
Skeena Valley The Skeena River is the second-longest river entirely within British Columbia, Canada (after the Fraser River). Since ancient times, the Skeena has been an important transportation artery, particularly for the Tsimshian and the Gitxsan—whose n ...
, overlapping with the range of
Sitka spruce ''Picea sitchensis'', the Sitka spruce, is a large, coniferous, evergreen tree growing to almost tall, with a trunk diameter at breast height that can exceed 5 m (16 ft). It is by far the largest species of spruce and the fifth-larg ...
(''Picea sitchensis''), and almost reaching the Arctic Ocean at latitude 69° N in the
District of Mackenzie The District of Mackenzie was a regional administrative district of Canada's Northwest Territories. The district consisted of the portion of the Northwest Territories directly north of British Columbia, Alberta, and Saskatchewan on Canada's main ...
, with white spruce up to 15 m high occurring on some of the islands in the Delta near
Inuvik Inuvik (''place of man'') is the only town in the Inuvik Region, and the third largest community in Canada's Northwest Territories. Located in what is sometimes called the Beaufort Delta Region, it serves as its administrative and service cen ...
. The wide variety of ecological conditions in which 4 Quebec conifers, including white spruce, are able to establish themselves, was noted by Lafond,Lafond, A. 1966. Notes sur l’écologie de quatre conifères du Québec: ''Picea mariana, P. glauca, Abies balsamea'', et ''Pinus banksiana''. Naturaliste Canadien, Québec 93:823–842. but white spruce was more exacting than
black spruce ''Picea mariana'', the black spruce, is a North American species of spruce tree in the pine family. It is widespread across Canada, found in all 10 provinces and all 3 territories. It is the official tree of the province of Newfoundland and Labra ...
. In the United States, the range of white spruce extends into Maine, Vermont, New Hampshire, New York, Michigan, Wisconsin, Minnesota, and Alaska,Sargent, C.S. 1922. Manual of the Trees of North America, 2nd corrected ed. Houghton and Mifflin, Boston, 510 p., reprinted 1961 in 2 volumes, Dover Publications, New York NY, Vol. 1. 433 p. 3999 bib gives 910 p./ref> where it reaches the Bering Strait in 66°44′ N" at Norton Bay and the Gulf of Alaska at Cook Inlet. Southern outliers have been reported in southern Saskatchewan and the Cypress Hills of southwestern SaskatchewanScoggan, H.J. 1957. Flora of Manitoba. Can. Dep. Northern Affairs and National Resources, Nat. Museum Can., Ottawa ON, Bull. 140. 619 p. and southeastern Alberta, northwestern Montana, south-central Montana, in the Black Hills on the Wyoming–South Dakota boundary, on the Manitoba–North Dakota boundary, and at Shushan, New York.Munns, E.N. 1938. The distribution of important forest trees of the United States. USDA, For. Serv., Washington DC, Misc. Publ. 287. 176 p.Fernald, M.L. 1950. Gray’s Manual of Botany, 8th ed. Amer. Book, New York NY. 1632 p.Cook, D.B.; Smith, R.H. 1959. A white spruce outlier at Shushan, New York. Ecology 40:333–337. White spruce is the northernmost tree species in North America, reaching just north of 69°N latitude in the Mackenzie River delta. It grows between sea level and an elevation of . Its northern distribution roughly correlates to the location of the tree line, which includes an isothermic value of for mean temperature in July, as well as the position of the Arctic front; cumulative summer degree days, mean net radiation, and the amount of light intensities also figure. White spruce is generally found in regions where the
growing season A season is a division of the year marked by changes in weather, ecology, and the amount of daylight. The growing season is that portion of the year in which local conditions (i.e. rainfall, temperature, daylight) permit normal plant growth. Whil ...
exceeds 60 days annually. The southern edge of the zone in which white spruce forms 60% or more of the total stand corresponds more or less to the July isotherm of around the Great Lakes; in the Prairie Provinces its limit is north of this isotherm. During the
summer solstice The summer solstice, also called the estival solstice or midsummer, occurs when one of Earth's poles has its maximum tilt toward the Sun. It happens twice yearly, once in each hemisphere ( Northern and Southern). For that hemisphere, the summer ...
, day length values range from 17 hours at its southern limits to 24 hours above the
Arctic Circle The Arctic Circle is one of the two polar circles, and the most northerly of the five major circles of latitude as shown on maps of Earth. Its southern equivalent is the Antarctic Circle. The Arctic Circle marks the southernmost latitude at w ...
. One of the hardiest conifers, white spruce in parts of its range withstands mean daily January temperature of and extreme minimum temperatures as low as ; minimum temperatures of are general throughout much of the range except the southernmost and southeasternmost parts.Fowells, H.A. 1965. ''Picea ''(spruces). p. 287–327'' in'' Silvics of Forest Trees of the United States. H.A. Fowells (Compiler), USDA, Forest Service, Washington DC, Agric. Handbook No. 271. By itself, or with black spruce and
tamarack ''Larix laricina'', commonly known as the tamarack, hackmatack, eastern larch, black larch, red larch, or American larch, is a species of larch native to Canada, from eastern Yukon and Inuvik, Northwest Territories east to Newfoundland, and als ...
(''Larix laricina''), white spruce forms the northern boundary of tree-form growth.Sutton, R.F. 1969. Silvics of white spruce (''Picea glauca'' oenchVoss). Can. Dep. Fish. For., For. Branch, Ottawa ON, Publ. 1250. 57 p. (Cited in Coates et al. 1994). White spruce up to 15 m in height occur at 69°N on islands in the Mackenzie Delta near Inuvik in the Northwest Territories. Hustich (1966)Hustich, I. 1966. On the forest–tundra and the northern tree-lines. Annales Univ. Turku A.II, Vol. 36:7–47. depicted ''Picea'' spp. as forming the northernmost limit of tree growth in North America. The arctic or northern timberline in North America forms a broad transition zone from Labrador to northern Alaska. In Labrador, white spruce is not abundant and constitutes less than 5% of the forest, with a range that coincides very closely with that of black spruce but extending slightly further north.Wilton, R.F. 1964. The forests of Labrador. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1066. 72 p. The range of white spruce extends westwards from Newfoundland and Labrador, and along the northern limit of trees to Hudson Bay, Northwest Territories, Yukon, and into northwestern Alaska. Across western Canada and Alaska, white spruce occurs further north than black spruce, and, while poplar (''Populus''), willow, and birch may occur along streams well into the tundra beyond the limits of spruce, the hardwoods are usually no more than scrub.Hustich, I. 1953. The boreal limits of conifers. Arctic 6:149–162. Spruce characteristically occurs in fingers of tree-form forest, extending far down the northern rivers and as scattered clumps of dwarfed “bush” spruce on intervening lands.Halliday, W.E.D.; Brown, A.W.A. 1943. The distribution of some important forest trees in Canada. Ecology 24:353–373. In Manitoba, Scoggan noted that the northernmost collection of white spruce was at latitude 59°48’N, but Bryson et al.Bryson, R.A.; Irving, W.H.; Larson, J.A. 1965. Radiocarbon and soil evidence of former forest in the southern Canadian tundra. Science 147(3653):46–48. found white spruce in the northern edge of continuous forest in central Canada at Ennadai Lake, about 60°45′ N, 101°’W, just north of the northwest corner of Manitoba. Bryson et al. noted that the forest retained “the same general characteristics as when it was first described Tyrell, J.B. 1897. Geol. Surv. Can., Ottawa ON, Ann. Rep. 1896, Vol. 9. (Cited by Bryson et al. 1965, orig. not seen)">y TyrrellTyrell, J.B. 1897. Geol. Surv. Can., Ottawa ON, Ann. Rep. 1896, Vol. 9. (Cited by Bryson et al. 1965, orig. not seen)in 1896”. Collins and Sumner reported finding white spruce within 13 km of the Arctic coast in the Firth valley, Yukon, at about 69°30′ N, 139°30′ W, and Sargent noted that white spruce in Alaska “reached Behring Strait in 66°44′ N”. Climate, especially temperature, is obviously a factor in determining distributions of northern flora. Halliday and Brown suggested that white spruce's northern limit corresponds “very closely” with the July mean monthly isotherm of 10 °C in Ungava, but that the northern limit west of Hudson Bay was south of that isotherm. Other climatic factors that have been suggested as affecting the northern limit of white spruce include: cumulative summer degree days, position of the Arctic front in July, mean net radiation especially during the growing season, and low light intensities. Topography, soil conditions, and glaciation may also be important in controlling northern limits of spruce.Drew, J.V.; Shanks, R.E. 1965. Landscape relationships of soils and vegetation in the forest–tundra ecotone, Upper Firth River Valley, Alaska–Canada. Ecol. Monogr. 35:285–306. The southern limit of distribution of white spruce is more complex. From east of the main range of coastal mountains in British Columbia, the southern continuous limit of white spruce is the forest/prairie interface through Alberta, Saskatchewan, Manitoba, the northern parts of Minnesota and Wisconsin, central Michigan, northeastern New York, and Maine. Sargent and Harlow and Harrar also included Vermont and New Hampshire; and, while Dame and BrooksDame, L.L.; Brooks, H. 1901. Handbook of the Trees of New England. Ginn, Boston MA. 196 p. excluded New York and states further west, they included Massachusetts as far south as Amherst and Northampton, “probably the southern limit of the species” in that area. NisbetNisbet, J. 1905. The Forester. Blackwood and Sons, Edinburgh and London, U.K., Vol. 1. 506 p. gave the range of white spruce as extending to “Carolina”, but he did not recognize red spruce as a species and presumably included it with white spruce. Towards the southern parts of its range, white spruce encounters increasingly effective ecological competition from
hardwood Hardwood is wood from dicot trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen. Hardwood (which comes from ...
s, some of which may reinforce their growth-rate or sprouting competitiveness with
allelopathic Allelopathy is a biological phenomenon by which an organism produces one or more biochemicals that influence the germination, growth, survival, and reproduction of other organisms. These biochemicals are known as allelochemicals and can have ben ...
depredation of coniferous
regeneration Regeneration may refer to: Science and technology * Regeneration (biology), the ability to recreate lost or damaged cells, tissues, organs and limbs * Regeneration (ecology), the ability of ecosystems to regenerate biomass, using photosynthesis ...
.Tubbs, C.H. 1976. Effect of sugar maple root exudate on seedlings of northern conifer species. USDA, For. Serv., Res. Note NC-213. 2 p. Further southward extension of the distribution is inhibited by white spruce's cold requirement.


White spruce as an exotic species

As an exotic, white spruce is widespread but uncommon. It was introduced into EnglandHereman, S. 1868. Paxton’s Botanical Dictionary (Revised and corrected), Bradbury, Evans, London, U.K. 623 p. and parts of continental EuropeJackson, A.B. 1948. The Identification of Conifers. Arnold, London, U.K. 152 p. in or soon after the year 1700, into Denmark about 1790,Sabroe, A.S. 1954. Forestry in Denmark, 3rd ed. Danish Heath Soc., Copenhagen. 118 p. and into Tasmania and
Ceylon Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
shortly before 1932.Troup, H.S. 1932. Exotic Forest Trees in the British Empire. Clarendon Press, Oxford, U.K. 268 p. Nisbet noted that firmly-rooted white spruce served very well to stabilize windswept edges of woods in Germany. In a narrow belt of mixed Norway and white spruces over an extremely exposed hilltop crest at high elevation in northern England, the Norway spruce were “completely dwarfed” whereas the white spruce had reached heights of between 3 and 4.3 m.Guillebaud, W.H.; Steven, H.M.; Marsden, R.E. 1920. Rate of growth of conifers in the British Isles. Forestry Commission, HMSO, London, U.K., Bull. 3. 84 p. The age of the belt was not recorded, but adjoining 66-year-old stands may have been of the same vintage. White spruce has also been used as a minor plantation species in England and Scotland.Selby, P.J. 1842. A history of British forest-trees. Van Voorst, London. 540 p.Anon. (C.P.J.) 1879. Fir. pp. 222–225 ''in'' vol IX. Encyclopædia Britannica, 9th ed.United Kingdom Forestry Commission. 1920. Beaufort estate. p. 57–62 ''in'' Programme, British Empire For. Conf., London, U.K. In Scotland, at Corrour, Inverness-shire, Sir John Stirling Maxwell in 1907 began using white spruce in his pioneering plantations at high elevations on deep peat. However, plantations in Britain have generally been unsatisfactory,Edlin, H.L. 1962. A modern sylva or a discourse of forest trees. 3. The spruces. Quart J. For. 56:292–300. mainly because of damage by spring frosts after mild weather had induced flushing earlier in the season. However, the species is held in high regard in the Belgian peat region, where it grows better than do the other spruces.Fraser, G.K. 1933. Studies of certain Scottish Moorlands in relation to tree growth. For. Commission, HMSO, London, U.K. 112 p.


Ecology

White spruce is a climax canopy tree in the boreal forests of Canada and Alaska. It generally occurs on well-drained soils in alluvial and riparian zones, although it also occurs in soils of glacial and lacustrine origin. The understory is dominated by feather mosses (''
Hylocomium splendens ''Hylocomium splendens'', commonly known as glittering woodmoss, splendid feather moss, stairstep moss, and mountain fern moss, is a Perennial plant, perennial Clone (plant), clonal moss with a widespread distribution in Northern Hemisphere borea ...
'', '' Pleurozium schreberi'', '' Ptilium crista-castrensis'') and fork mosses, and occasionally peat moss. In the far north, the total depth of the moss and underlying
humus In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
is normally between , although it tends to be shallower when
hardwood Hardwood is wood from dicot trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen. Hardwood (which comes from ...
s are present in the stand. White spruce grows in soils with pH values of 4.7–7.0, although they have been found in soils as acidic as 4.0 in subalpine fir forests in the Northwest Territories. A presence of calcium in the soil is common to white spruce found in northern New York. White spruce most commonly grows in the soil orders of
Alfisols Alfisols are a soil order in USDA soil taxonomy. Alfisols form in semi-arid to humid areas, typically under a hardwood forest cover. They have a clay-enriched subsoil and relatively high native fertility. "Alf" refers to aluminium (Al) and iron ( ...
and Inceptisols. Soil properties such as fertility, temperature, and structural stability are partial determinants of the ability of white spruce to grow in the extreme northern latitudes. In the northern limits of its range, white spruce is the climax species along with black spruce; birch and aspen are the early succession species. Wildfires typically occur every 60 to 200 years, although they have been known to occur as infrequently as every 300 years. White spruce will grow in USDA Growing Zones 3–7, but is not adapted to heat and humidity and will perform poorly in a hot climate. The tree attains its greatest longevity and growth potential in Zones 3–4.


Soils

White spruce occurs on a wide variety of soils, including soils of glacial, lacustrine, marine, and alluvial origins; overlying basic dolomites, limestones and acidic Precambrian and Devonian granites and gneisses; and Silurian sedimentary schists, shales, slates, and conglomerates.Halliday, W.E.D. (1937). "A forest classification for Canada." Can. Dep. Mines and Resources, Dominion For. Serv., Ottawa ON, Bull. 89. 50 p. The wide range of textures accommodated includes clays, even those that are massive when wet and columnar when dry, sand flats, and coarse soils.Wilde, S.A.; Wilson, F.G.; White, D.P. (1949). "Soils of Wisconsin in relation to silviculture." Wisconsin Conserv. Dep., Madison WI, Publ. 525–49. 171 p.Wilde, S.A.; Voigt, G.K.; Pierce, R.S. (1954). "The relationship of soils and forest growth in the Algoma district of Ontario, Canada." ''J. Soil Sci.'' 5:22–38.Nienstaedt, H. (1957). Silvical characteristics of white spruce (''Picea glauca''). USDA, For. Serv., Lake States For. Exp. Sta., St. Paul MN, Pap. 55. 24 p.Jameson, J.S. (1963). "Comparison of tree growth on two sites in the Riding Mountain Forest Experimental Area." Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1019. 36 p. Its occurrence on some organic soils is not characteristic, except perhaps on shallow mesic organic soils in Saskatchewan and in association with black spruce on organic soils in central Yukon. Podzolized, brunisolic, luvisolic, gleysolic, and regosolic (immature) soils are typical of those supporting white spruce throughout the range of the species. Soils supporting white spruce are most commonly Alfisols or Inceptisols. In the podzol region of Wisconsin, white spruce occurs on
loam Loam (in geology and soil science) is soil composed mostly of sand (particle size > ), silt (particle size > ), and a smaller amount of clay (particle size < ). By weight, its mineral composition is about 40–40–20% concentration of sand–sil ...
podzols, podzolized gley loams, strongly podzolized clays, gley-podzol clays, stream-bottom soils, and wood peat. Moist sandy loams also support good growth. On sandy podzols, it is usually a minor species. Good development occurs on moist alluvium on the banks of streams and borders of swamps.Jeffrey, W.W. (1961). "Origin and structure of some white spruce stands on the lower Peace River." Can. Dep. For., For. Res. Branch, Ottawa ON, Tech. Note 103. 20 p.Jeffrey, W.W. (1964). "Forest types along lower Liard River, Northwest Territories." Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1035. 103 p.Lacate, D.S.; Horton, K.W.; Blyth, A.W. (1965). "Forest conditions on the Lower Peace River." Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1094. 53 p.Viereck, L.A. (1973). "Wildfire in the taiga of Alaska." ''Quaternary Res.'' 3:465–495.Sargent, C.S. 1898. The Silva of North America. A description of the trees which grow naturally in North America exclusive of Mexico. Vol. XII. Coniferae. Houghton Mifflin, Riverside Press, Cambridge, Boston MA. 144 p.Kenety, W.H. (1917). "Preliminary study of white spruce in Minnesota." ''Univ. Minnesota, Cloquet Exp. Sta. MN, Bull.'' 168. 30 p. White spruce makes good growth on well-drained lacustrine soils in Alberta Mixedwoods,Heger, L. (1971). "Site-index/soil relationships for white spruce in Alberta mixedwoods." Can. Dep. Environ., Can. For. Serv., For. Manage. Instit., Ottawa ON, Inf. Rep. FMR-X-32. 15 p. on moderately-well-drained clay loams in Saskatchewan,,Kabzems, A. (1971). "Growth and yield of well stocked white spruce in the mixedwood section , Saskatchewan. Saskatchewan." Dep. Nat. Resour., For. Branch, Prince Albert SK, Tech. Bull. 5. 75 p. (Cited in Coates et al. 1994). and on melanized loams and clays (with sparse litter and a dark-coloured organically-enriched mineral horizon) in the Algoma district of Ontario. White spruce becomes less accommodating of soil with increasing severity of climate. The distribution of white spruce in Labrador seems to depend almost entirely on the character of the soil, and between the southwestern shores of Hudson Bay and the northeastern regions of Saskatchewan, white spruce is confined to very local physiographic features, characterized by well-drained or fertile soils.Ritchie, J.C. 1956. The vegetation of northern Manitoba. I. Studies in the southern spruce forest zone. Can. J. Bot. 34(4):523–561. On dry, deep, outwash deposits in northern Ontario, both white spruce and aspen grow slowly.MacLean, D.W. (1960). "Some aspects of the aspen–birch–spruce–fir type in Ontario." Can. Dep. Northern Affairs National Resources, For. Branch, For. Res. Div., Ottawa ON, Tech. Note 94. 24 p. But, broadly, white spruce is able to tolerate considerable droughtiness of sites that are fertile, and no fertile site is too moist unless soil moisture is stagnant.Sutton, R.F. (1968). Ecology of young white spruce (''Picea glauca'' oenchVoss). Ph.D. thesis, Cornell Univ., Ithaca NY, Univ. Microfilms, Ann Arbor, Michigan MI, 68–11645. 500 p. Soil fertility holds the key not just to white spruce growth but to the distribution of the species. At least moderate fertility is needed for good growth, but white spruce occurs on many sites where nutrient deficiencies depress its growth more than that of black spruce, red spruce, Norway spruce, and the pines generally. Minimum soil-fertility standards recommended for white spruce sufficient to produce 126 to 157 m3/ha of wood at 40 years are much higher than for pine species commonly planted in the Lake States (Wilde 1966):Wilde, S.A. (1966). "Soil standards for planting Wisconsin conifers." ''J. For.'' 64(6):389–391. 3.5% organic matter, 12.0 meq/100 g exchange capacity, 0.12% total N, 44.8 kg/ha available P, 145.7 kg/ha available K, 3.00 meq/100 g exchangeable Ca, and 0.70 meq/100 g exchangeable Mg. Forest floors under stands dominated by white spruce respond in ways that vary with site conditions, including the disturbance history of the site. Composition,
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
, and mineral soil physical and chemical properties are affected. In Alaska, the accumulation of organic layers (to greater thicknesses in mature stands of spruce than those in hardwood stands on similar sites) leads to decreased soil temperatures, in some cases leading to the development of permafrost.Viereck, L.A. (1970a). "Forest succession and soil development adjacent to the Chena River in interior Alaska." ''Arctic Alp. Res.'' 2(1):1–26. S. BA51:76183/ref>Viereck, L.A. (1970b). Soil temperatures in river bottom stands in interior Alaska. p. 223–233 ''in'' Proc. Ecology of the Subarctic Regions, July–Aug. 1966, Helsinki, Finland, UNESCO. ienstaedt and Zasada 1990/ref>Viereck, L.A.; Dyrness, C.T.; Van Cleve, K.; Foote, M.J. (1983). "Vegetation, soils, and forest productivity in selected forest types in interior Alaska." ''Can. J. For. Res.'' 13(5):703–720. Acidity of the mineral soil sampled at an average depth of 17 cm in 13 white spruce stands on abandoned farmland in Ontario increased by 1.2 pH units over a period of 46 years.Brand, D.G.; Kehoe, P.; Connors, M. (1986). "
Coniferous Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extant ...
afforestation leads to soil acidification in central Ontario." ''Can. J. For. Res.'' 16(6):1389–1391.
A considerable range of soil pH is tolerated by white spruce. Thrifty stands of white spruce in Manitoba have developed on soils of pH 7.6 at only 10 cm below the surface, and pH 8.4 at 43 cm below the surface;Stoeckeler, J.H. (1938). "Soil adaptability of white spruce." ''J. For.'' 36:1145–1147. rooting depth in those soils was at least 81 cm. An abundant calcium supply is common to most white spruce locations in New York state. Chlorosis was observed in young white spruce in heavily limed nursery soils at about pH 8.3. Wilde gave 4.7 to 6.5 as the approximate optimum range of pH for white spruce in Wisconsin, but optimum growth seems possible at pH levels up to 7.0 and perhaps higher. Alluvium on the floodplains of northern rivers shows pH levels from 5.0 to 8.2.Zasada, J.C.; Van Cleve, K.; Werner, R.A.; McQueen, J.A.; Nyland, E. (1977). "Forest biology and management in high-latitude North American forests." p. 137–195'' in'' Proc. North American Forest Lands at Latitudes North of 60 degrees. Sympos., Univ. Alaska, Fairbanks AK, Sept. 19–22, 1977. High-lime ecotypes may exist,Pelletier, J.R. (1966). "Tree breeding in Canada." ''Commonw. For. Rev.'' 45(1):9–10. and in Canada Forest Section B8 the presence of balsam poplar and white spruce on some of the moulded moraines and clays seems to be correlated with the considerable lime content of these materials,Stiell, W.M. (1976). "White spruce: artificial regeneration in Canada." Dep. Environ., Can. For. Serv., Ottawa ON, Inf. Rep. FMR-X-85. 275 p. while calcareous soils are favourable sites for northern outliers of white spruce. Mature stands of white spruce in boreal regions often have well-developed
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
layers dominated by feather mosses, e.g., ''
Hylocomium splendens ''Hylocomium splendens'', commonly known as glittering woodmoss, splendid feather moss, stairstep moss, and mountain fern moss, is a Perennial plant, perennial Clone (plant), clonal moss with a widespread distribution in Northern Hemisphere borea ...
'', '' Pleurozium schreberi'', '' Ptlium crista-castrensis'', and ''
Dicranum ''Dicranum'' is a genus of mosses, also called wind-blown mosses or fork mosses. These mosses form in densely packed clumps. Stems may fork, but do not branch. In general, upright stems will be single but packed together. ''Dicranum'' is distrib ...
'', rather than '' Sphagnum''.La Roi, G.H.; Stringer, M.H. (1976). Ecological studies in the boreal spruce–fir forests of the North American taiga. II. Analysis of the bryophyte flora. Can. J. Bot. 54:619–643. ienstaedt and Zasada 1990/ref>Viereck, E.G. (1987). Alaska’s wilderness medicines – healthful plants of the North. Alaska Publishing, Edmonds, Washington WA. 107 p. ienstaedt and Zasada 1990/ref> The thickness of the moss–organic layer commonly exceeds 25 cm in the far north and may approach twice that figure. The mosses compete for nutrients and have a major influence on soil temperatures in the rooting zone. Permafrost development in parts of Alaska, Yukon, and the Northwest Territories is facilitated by the insulative organic layer (Viereck 1970a, b, Gill 1975, Van Cleve and Yarie 1986).Gill, D. (1975). "Influence of white spruce trees on permafrost-table microtopography, Mackenzie River Delta." ''Can. J. Earth Sci.'' 12(2):263–272.Van Cleve, K.; Yarie, J. (1986). "Interaction of temperature, moisture, and soil chemistry in controlling nutrient cycling and ecosystem development in the taiga of Alaska." p. 160–189 ''in'' Van Cleve, K.; Chapin, F.S.; Flanagan, P.W.; Viereck, L.A.; Dyrness, C.T. (Eds.). (1986). ''Forest Ecosystems in the Alaskan Taiga.'' Springer-Verlag, New York NY.


Cold hardiness

White spruce is extremely hardy to low temperatures, provided the plant is in a state of winter
dormancy Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be clo ...
. Throughout the greater part of its range, white spruce routinely survives and is undamaged by winter temperatures of , and even lower temperatures occur in parts of the range.
Boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
''Picea'' are among the few extremely hardy conifers in which the bud primordia are able to survive temperatures down to .Sakai, A.; Larcher, W. (Eds.) (1987). ''Frost Survival of Plants.'' Springer-Verlag, New York NY. 321 p. Especially important in determining the response of white spruce to low temperatures is the physiological state of the various tissues, notably the degree of "hardening" or dormancy. A natural progression of hardening and dehardening occurs in concert with the seasons.Glerum, C. (1985). "Frost hardiness of coniferous seedlings: principles and applications." p. 107–123 ''in ''Duryea, M.L. (Ed.). Proceedings: Evaluating seedling quality: principles, procedures, and predictive abilities of major tests. Workshop, October 1984, Oregon State Univ., For. Res. Lab., Corvallis OR. While different tissues vary in ability to tolerate exposure to stressful temperatures, white spruce, as with woody plants in general, has necessarily developed sufficient winter hardiness in its various tissues to enable them to survive the minimum temperatures experienced in the distribution range. White spruce is subject to severe damage from spring frosts. Newly flushed shoots of white spruce are very sensitive to spring frost.Smith, B.J. (1949). "Silvicultural work at the Sault Ste. Marie Division
f Abitibi Power and Paper Co. Ltd. F, or f, is the sixth letter in the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages and others worldwide. Its name in English is ''ef'' (pronounced ), and the plural is ''efs''. Hist ...
" Can. Pulp Paper Assoc., Woodlands Section, Woodlands Section Index No. 1050 (F-2). 4 p.
This sensitivity is a major constraint affecting young trees planted without overstorey nurses in boreal climates.Sutton, R.F. (1992). "White spruce (''Picea glauca'' oenchVoss): stagnating boreal old-field plantations unresponsive to fertilization and weed control." ''For. Chron.'' 68:249–258.


Forest succession

Forest succession in its traditional sense implies two important features that resist direct examination.Solomon, A.M.; West, D.C., and Solomon, J.A. 1981. Simulating the role of climate change and species imiigration in forest succession. p. 154–178 ''in'' West, D.C.; Shugart, H.H.; Botkin, D.B. (Eds.). Forest Succession: Concepts and Application. Springer-Verlag, New York NY. First, classical definitions generally connote directional changes in species composition and community structure through time, yet the time frame needed for documentation of change far exceeds an average lifespan. The second feature that defies quantitative description is the end point or climax. Floodplain deposits in the Northwest Territory, Canada, are important in relation to the development of productive forest types with a component of white spruce. The most recently exposed surfaces are occupied by sandbar vegetation or riparian shrub willows and '' Alnus incana''. With increasing elevation, the shrubs give way successively to balsam poplar and white spruce forest. In contrast, older floodplains, with predominantly brown wooded soils, typically carry white spruce– trembling aspen mixedwood forest. Interrelationships among nutrient cycling, regeneration, and subsequent forest development on floodplains in interior Alaska were addressed by Van Cleve et al.,Van Cleve, K.; Dyrness, R.; Viereck, L. 1980. Nutrient cycling in interior Alaska flood plains. p. 11–18 ''in'' Murray, M.; Van Veldhuizen, R.M. (Eds.). Proc. Workshop, Fairbanks, Alaska, Nov. 1979. USDA, For. Serv., Pacific Northwest For. Exp. Sta., Portland OR, Gen. Tech. Rep., PNW-107. 52 p. who pointed out that the various stages in primary succession reflect physical, chemical, and biological controls of ecosystem structure and function. Thus, each successional stage has a species combination in harmony with site quality. Short-circuiting succession by planting a late successional species such as white spruce on an early successional surface may result in markedly reduced growth rates because of nitrogen insufficiency. Without application of substantial amounts of fertilizer, use would have to be made of early successional alder and its site-ameliorating additions of nitrogen. Neiland and Viereck noted that “the slow establishment and growth of spruce under birch stands
n Alaska N, or n, is the fourteenth letter in the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages and others worldwide. Its name in English is ''en'' (pronounced ), plural ''ens''. History ...
may be partially due to effects of shading and general competition for water and nutrients, but may also be more directly related to the birch itself. HeikinheimoHeikinheimo, O. 1915. Der einfluss der brandwirtschaft auf die Wälder Finnlands. Kaskiviljelyksen Vaikutus Suomen Metsin. Acta Forest. Fenn. 4:1–264, 1–149, 1–59 erman summary p 1–59/ref>Lutz, H.J. 1956. Ecological effects of forest fires in the interior of Alaska. USDA, For. Serv., Washington DC, Tech. Bull. 1133. 121 p. found that birch ash inhibited white spruce seedlings, and GregoryGregory, R.A. 1966. The effect of leaf litter upon establishment of white spruce beneath paper birch. For. Chron. 42:251–255. found that birch litter has a smothering effect on spruce seedlings.".Neiland, B.J.; Viereck, L.A. 1977. Forest types and ecosystems. p. 109–136 ''in'' North American Forest Lands at Latitudes North of 60 Degrees, Proc. sympos., Univ. Alaska, Fairbanks AK, Sept. 1977. On dry upland sites, especially south-facing slopes, the mature vegetation is white spruce, white birch, trembling aspen, or a combination of these species. Succession follows in one of two general patterns. In most cases, aspen and birch develop as a successional stage after fire before reaching the spruce stage. But, occasionally, with optimal site conditions and a source of seed, white spruce will invade with the
hardwood Hardwood is wood from dicot trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen. Hardwood (which comes from ...
s or within a few years thereafter, thereby producing even-aged white spruce stands without an intervening hardwood stage.


Associated forest cover

The White Spruce Cover Type may include other species in small numbers. In Alaska, associates include paper birch, trembling aspen, balsam poplar, and black spruce; in western Canada, additional associates are subalpine fir, balsam fir, Douglas-fir, jack pine, and lodgepole pine.Dyrness, C.T. 1980. Western forest cover types, Northern Interior (Boreal): White spruce. p.81; White spruce–Aspen. p.82; Black spruce–White spruce. p.84; and Black spruce–Paper birch. p.85. ''in'' Eyre, F.H. (Ed.). Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC. Seral species giving way to white spruce include paper birch, aspen, balsam poplar, jack pine, and lodgepole pine. On certain river bottom sites, however, black spruce may replace white spruce. Earlier successional stages leading to the white spruce climax are the white spruce–paper birch, white spruce–aspen, balsam poplar, jack pine, and lodgepole pine types. The type shows little variation. The forest is generally closed and the trees well formed, other than those close to the timberline. Lesser vegetation in mature stands is dominated by mosses. Vascular plants are typically few, but
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
s and
herb In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal ...
s that occur “with a degree of regularity” include: alder, willows, mountain cranberry, red-fruit bearberry, black crowberry, prickly rose, currant, buffaloberry, blueberry species, bunchberry, twinflower, tall lungwort, northern comandra, horsetail, bluejoint grass, sedge species, as well as ground-dwelling mosses and
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship. Of the Eastern Forest Cover Types recognized by the Society of American Foresters, only one, White Spruce, names that species in its title. The eastern White Spruce Cover Type, as defined, encompasses white spruce both in pure stands, and in mixed stands "in which white spruce is the major ndefinedcomponent."Payette, S. 1980. Eastern forest cover types, Boreal Forest Region: white spruce. p.15 ''in'' Eyre, F.H. (Ed.). Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC. 148 p. In most of its range, white spruce occurs more typically in association with trees of other species than in pure stands. White spruce is an associated species in the following Eastern Forest cover types, by the Society of American Foresters; in the Boreal Forest Region: (1) jack pine, (5) balsam fir, (12) black spruce, (16) aspen, (18) paper birch, and (38) tamarack; in the Northern Forest Region: (15) red pine, (21) eastern white pine, (24) hemlock-yellow birch, (25) sugar maple-beech-yellow birch, (27) sugar maple, (30) red spruce-yellow birch, (32) red spruce, (33) red spruce-balsam fir, (37) northern white-cedar, and (39) black ash-American elm-red maple.Eyre, F.H. (Ed.) 1980. Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC. 148 p.


Predators

Outbreaks of spruce beetles have destroyed over of forests in Alaska. Although sometimes described, e.g., by Switzer (1960),Switzer, A.L.K. (1960). "Spruce management for the future." ''For. Chron.'' 36(2):163–165. as relatively resistant to attack by insects and disease, white spruce is far from immune to depredation. Important insect pests of white spruce include the spruce budworm (''Choristoneura fumiferana''), the yellow-headed spruce sawfly (''
Pikonema alaskensis ''Pikonema'' is a genus of sawflies belonging to the family Tenthredinidae Tenthredinidae is the largest family of sawflies, with well over 7,500 species worldwide, divided into 430 genera. Larvae are herbivores and typically feed on the foli ...
''), the European spruce sawfly (''
Gilpinia hercyniae ''Gilpinia hercyniae'', the European spruce sawfly, was first found in North America near Hull, Quebec, in 1922 (Rose and Lindquist 1985).Rose, A.H.; Lindquist, O.H. 1985. Insects of eastern spruces, fir and, hemlock, revised edition. Gov’t Ca ...
''), the spruce bud moth (''
Zeiraphera canadensis ''Zeiraphera canadensis'', the spruce bud moth, is a moth of the family Tortricidae. It is a small brown moth mainly found in North America, specifically New Brunswick, Quebec, and the north-eastern United States. The adult moth flutters quickly ...
''), and
spruce beetle ''Dendroctonus rufipennis'', the spruce beetle, is a species of bark beetle native to British Columbia, Newfoundland and Labrador, Nova Scotia, Ontario, Quebec, Northern Manitoba, the Yukon, Colorado, Wyoming, Montana, and Maine. They are known ...
(''Dendroctonus rufipennis'').Fowells, H.A. (1965). ''Picea'' (spruces). p. 287–327 in ''Silvics of Forest Trees of the United States.'' H.A. Fowells (Compiler), USDA, Forest Service, Washington DC, Agric. Handbook No. 271.Rose, A.H.; Lindquist, O.H. (1985). ''Insects of eastern spruces, fir and, hemlock, revised edition.'' Gov’t Can., Can. For. Serv., Ottawa, For. Tech. Rep. 23. 159 p. (cited in Coates et al. 1994, cited orig ed 1977)Ives, W.G.H.; Wong, H.R. (1988). ''Tree and shrub insects of the prairie provinces.'' Gov’t Can., Can. For. Serv., Edmonton AB, Inf. Rep. NOR-X-292. 327 p.
oates et al. 1994 Oates may refer to: ;People * Oates (surname) *Garfunkel and Oates, comedy act ** Kate Micucci (aka ''Oates'') of "Garfunkel and Oates" *Hall & Oates, musical act ;Places and buildings * Oates, Missouri, a community in the United States *Oates Coa ...
/ref> As well, other budworms, sawflies, and bark beetles, gall formers, bud midges, leaf miners, aphids, leaf eaters, leaf rollers, loopers, mites, scales, weevils, borers, pitch moths, and spittlebugs cause varying degrees of damage to white spruce. A number of sawflies feed on spruce trees. Among them European spruce sawfly, yellow-headed spruce sawfly, green-headed spruce sawfly and the spruce webspinning sawfly. More than a dozen kinds of looper feed on the spruces, fir, and hemlock in eastern Canada. The full-grown larvae of the larvae vary in length from 15 mm to 35 mm. Some feed briefly in the fall and complete their feeding in the spring; others feed mainly in the summer; still others feed mainly in the late summer and fall. The fall and spring feeding group includes the dash-lined looper (''Protoboarmia porcelaria indicataria''), the diamond-backed looper (''Hypagyrtis piniata''), the fringed looper (''Campaea perlata''), and the false loopers (''Syngrapha'' species). The summer feeding group includes the false hemlock looper (''Nepytia canosaria'' Walker), occasionally occurring in large numbers and usually in conjunction with the
hemlock looper ''Lambdina fiscellaria'', the mournful thorn or hemlock looper, is a moth of the family Geometridae. It is found in North America, from the Pacific to the Atlantic coast and from Canada south to Pennsylvania, Wisconsin and California. The adult ...
(''Lambdina fiscellaria''), the '' Eupithecia'' species, the yellowlined conifer looper (''Cladara limitaria''), and the saddleback looper (''Ectropis crepuscularia''). The late summer and fall group includes the common spruce-fir looper (''Semiothisa signaria dispuncta'') and the similar
hemlock angle ''Macaria fissinotata'', the hemlock angle, is a moth of the family Geometridae. It is found in North America from Nova Scotia to Georgia, west to Kentucky, north to Ontario. The wingspan is 22–25 mm. The moth flies from May to Septemb ...
(moth) ''Macaria fissinotata'' on hemlock, the small spruce loopers '' Eupithecia'' species, the
gray spruce looper ''Caripeta divisata'', known generally as the gray spruce looper or twin-spot girdle, is a species of geometrid moth in the family Geometridae. It is found in North America. The MONA or Hodges number for ''Caripeta divisata'' is 6863. Refere ...
''Caripeta divisata'', occasionally abundant, the
black-dashed hydriomena moth ''Hydriomena divisaria'', the black-dashed hydriomena moth, is a moth in the family Geometridae. It is found in North America, where it has been recorded from southern Canada and the northern United States, south to Georgia in the east. The wi ...
(''Hydriomena divisaria''), and the
whitelined looper ''Epirrita pulchraria'', the whitelined looper, is a species of geometrid moth in the family Geometridae. The MONA or Hodges number for ''Epirrita pulchraria'' is 7435. References Further reading * * Epirrita Articles created by Qb ...
(''Eufidonia notataria'').


Uses

Wildlife such as deer, rabbits, and grouse browse the foliage during the winter. The wood of white spruce is of a lower quality than that of Engelmann spruce, but is stronger. It was used to make shelters and as firewood by Native Americans and European settlers in Alaska, where lodgepole pine does not grow. The wood is of major economic importance in Canada, being harvested for paper and construction. It is also used as a
Christmas tree A Christmas tree is a decorated tree, usually an evergreen conifer, such as a spruce, pine or fir, or an artificial tree of similar appearance, associated with the celebration of Christmas. The custom was further developed in early modern ...
. The wood is also exported to
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
where, known as "shin-kaya", it is used to make go boards as a substitute for the rare
kaya Kaya may refer to: People *Kaya (given name) *Kaya (surname) Places *Kaya, Burkina Faso, a town in Burkina Faso, capital of the department *Kaya Airport, serving the town * Kaya Department, a department or commune of Sanmatenga Province in centr ...
wood. Black Hills Spruce (''Picea glauca'' var. ''densata'') is used for
bonsai Bonsai ( ja, 盆栽, , tray planting, ) is the Japanese art of growing and training miniature trees in pots, developed from the traditional Chinese art form of ''penjing''. Unlike ''penjing'', which utilizes traditional techniques to produce ...
. White spruce is the provincial tree of Manitoba and the
state tree This is a list of U.S. state, federal district, and territory trees, including official trees of the following of the states, of the federal district, and of the territories. Table See also * List of U.S. state, district, and territorial insign ...
of South Dakota. The new growth or tips of white spruce is used in beer making, gin production, flavouring pop, candy making or in pickles and preserves.


Varieties

Several geographical varieties have been described, but are not accepted as distinct by all authors. These comprise, from east to west: *''Picea glauca'' var. ''glauca'' (typical or eastern white spruce): from Newfoundland west to eastern Alberta, on lowland plains. *''Picea glauca'' var. ''densata'' (Black Hills white spruce): The Black Hills in South Dakota. *''Picea glauca'' var. ''albertiana'' (Alberta white spruce): The Rocky Mountains in Alberta, British Columbia and northwest Montana. *''Picea glauca'' var. ''porsildii'' (Alaska white spruce): Alaska and Yukon. The two western varieties are distinguished by pubescent shoots, and may be related to extensive hybridisation and intergradation with the closely related Engelmann spruce found further south in the Rocky Mountains. White spruce also hybridises readily with the closely related Sitka spruce where they meet in southern Alaska and northwestern British Columbia; this hybrid is known as ''
Picea × lutzii ''Picea'' × ''lutzii'' is a hybrid spruce tree that is a natural cross between white spruce and Sitka spruce occurring where the ranges of the two species overlap in coastal south-central Alaska and coastal British Columbia. Its common name is L ...
''.


Cultivars

Numerous
cultivars A cultivar is a type of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and when Plant propagation, propagated retain those traits. Methods used to propagate cultivars include: division, root and st ...
of various sizes, colours and shapes have been selected for use in parks and gardens. The following have gained the Royal Horticultural Society's
Award of Garden Merit The Award of Garden Merit (AGM) is a long-established annual award for plants by the British Royal Horticultural Society (RHS). It is based on assessment of the plants' performance under UK growing conditions. History The Award of Garden Merit ...
. *''Picea glauca'' ‘Echiniformis’ *''Picea glauca ''var.'' albertiana'' ‘Alberta Globe’ *''Picea glauca ''var.'' albertiana'' ‘Conica’ 'Conica' is a dwarf conifer with very slender leaves, like those normally found only on one-year-old seedlings, and very slow growth, typically only per year. Older specimens commonly ' revert', developing normal adult foliage and starting to grow much faster; this 'reverted' growth must be
pruned Pruning is a horticultural, arboricultural, and silvicultural practice involving the selective removal of certain parts of a plant, such as branches, buds, or roots. The practice entails the ''targeted'' removal of diseased, damaged, dead, ...
if the plant is to be kept
dwarf Dwarf or dwarves may refer to: Common uses *Dwarf (folklore), a being from Germanic mythology and folklore * Dwarf, a person or animal with dwarfism Arts, entertainment, and media Fictional entities * Dwarf (''Dungeons & Dragons''), a humanoid ...
.


Genome

''P. glauca'' has three different genomes; a nuclear genome, a mitochondrial genome, and a plastid (i.e. chloroplast) genome. The large (20 Gbp) nuclear genome of ''P. glauca'' (genotype WS77111) was published in 2015, and the organellar (plastid and mitochondrial) genomes (genotype PG29) were published in SD Jackman et al. 2015. The plastid genome of ''P. glauca'' (genotype WS77111) has also been published.


References


External links

* {{Authority control Least concern plants Ornamental trees glauca Provincial symbols of Manitoba Taiga and boreal forests Trees of humid continental climate Trees of continental subarctic climate Trees of Alberta Trees of British Columbia Trees of Canada Trees of Manitoba Trees of Ontario Trees of Saskatchewan Trees of Western Canada Trees of Subarctic America Trees of Eastern Canada Trees of Alaska Trees of the Northeastern United States Trees of the Northwestern United States Trees of the North-Central United States Trees of the United States Trees of the Great Lakes region (North America) Trees of North America Symbols of South Dakota